directional selection is when the population evolves toward the fittest genotype, with the allele that offer the highest fitness becoming more common over gens until it reaches fixation
if the fitness of heterozygote is exactly between the fitness of the two homozygotes, the population will move towards the fittest genotype (homozygote) over generatrions
1. inital allele frequencies
2. selection coeffcient
3. generation time
4. degree of dominance
5. population size
rare alleles: take longer to reach fization as they start with low frequency
common alleles: reach fixation due to faster high intial frequency
a larger selection coefficient (s) means a larger fitness advantage, which causes the beenfical allele to rise to fixation more quickly
species with shorter gen times will see beneficial alleles reach fixation faster than species with longer gen times which take longer to evolve
completely dominant allles rise to fixation fatest since they are expressed in both heterozygotes and homzygotes where as recessive alleles rise slowly at first but then rapidly progress to fixation after reaching a certain frequency
they are slower at first than complete dominant but reach the fixation the fatest becuase both homozygotes express the trait
small pops are more affected by genetic drift which can cause beneficial alleles which are present at a low frequency to be lost by drift/chance because they wont increase in freq
large pops experience less drift, allowing selection to operate more effciently and benefical alleles to reach fixation faster
as the frequency of either allele A1 or A2 increases, its relative fitness decreases. this means the alllele becomes less fit as it becomes more common in the pop
When the frequency of an allele increases, its relative fitness decreases, which leads to a decrease in the allele’s frequency in the population over time. This is a feature of negative frequency-dependent selection.
Negative frequency-dependent selection occurs when a genotype has increased fitness only when it is rare in the population. As its frequency increases, its fitness decreases, leading to a balancing effect where the rare genotype is favored by selection.
Yes, negative frequency-dependent selection is actually quite common in nature. It helps maintain genetic diversity by favoring rare genotypes.
An example is the elder flower orchid (Dactylorhiza sambucina) from Europe, which has two color morphs: purple and yellow. The orchid doesn't contain nectar, so pollinators aren't rewarded. However, the rarer color morph has higher reproductive success.
As the yellow morph of the elder flower orchid becomes more common, its reproductive success decreases, which is an example of negative frequency-dependent selection. The rare color morph is always favored, and its frequency increases.
Over time, the population swings back and forth between the two color morphs, with selection pressure always favoring the rarer color. This results in a balance being maintained between the two color morphs in the population.
The rarer color morph (either purple or yellow) is favored, leading to fluctuating population frequencies of each color. The mean frequency of both morphs is maintained over time due to the selection pressure.
Fitness is calculated as absolute fitness, which is the product of the number of offspring and the survival rate of an individual:
Fitness=Offspring×Survival Rate
Relative fitness compares an individual's fitness to the most fit individual in the population. The most fit individual is assigned a relative fitness of 1. To calculate relative fitness, use the following:
Relative Fitness (w)=
Absolute Fitness of Genotype
/Highest Absolute Fitness
Mean fitness is the average fitness of all individuals in the population. It can be calculated by taking the weighted average of the relative fitness of all genotypes, where the most fit genotype has a relative fitness of 1.
Delta p represents the change in allele frequency between generations. If,
Δp>0, it indicates that the frequency of a particular allele is increasing in the population. postive sslection, genetic drift in small pops, mutations, gene flow, non random mating.
Δp=. 0 allele freq is constant, no evolutionary forces acting upon allele.
Δp<0 allle freq dec, maybe cause of natural selection, genetic drift, or gene flow
Pleiotropy is when a single gene influences multiple traits or phenotypic characteristics. A mutation in a pleiotropic gene can have wide-ranging effects on an organism.
Most traits are distributed on a bell curve because they are polygenic (controlled by multiple genes) and influenced by the environment. This results in a normal distribution, with phenotypes clustered around a mean value.
The total variance (Vp) is the sum of environmental variance (Ve) and genetic variance (Vg): Vp=Ve+Vg
Variation due to environmental influences
Vi: Interactive variance (genes interact with each other)
Va: Additive variation (multiple genes contribute additively to the phenotype)
Vd: Dominance variation (due to dominant-recessive allele interactions)
H² : Broad sense heritability represents the proportion of the total variance in a trait due to genetic factors (both additive, dominance, and interaction).
h²: Narrow sense heritability measures the proportion of variance in a trait due to additive genetic effects only (i.e., the alleles that contribute to the trait in a straightforward, additive way).
If H² is high, it indicates the trait is more heritable (closer to 1).
Balancing Selection, Disruptive Selection, Directional Selection, Stabilizing Selection
Maintains genetic diversity by favoring the maintenance of multiple alleles.
Selects for extreme phenotypes, causing an increase in variance and potentially leading to a bimodal distribution.
Favors one extreme phenotype, causing a shift in the mean of the population and possibly reducing variance.
Selects for average phenotypes, reducing variance but keeping the mean unchanged.
In stabilizing selection, the mean remains the same, but variance decreases as selection favors the average phenotype and eliminates extreme variants.
In disruptive selection, the mean remains unchanged, but variance increases as the population is selected against the average phenotype and favors the extremes, leading to a bimodal distribution.
In directional selection, the mean changes in favor of one extreme, while the variance remains relatively the same. This leads to a shift toward the favored extreme phenotype.
The response to selection depends on heritability—the higher the heritability ( h²) the faster the response to selection. Traits with high heritability are more likely to show rapid changes in response to selective pressure.
Big S represents the strength of selection, which is the difference between the mean trait value in the population and the mean trait value in the reproducing individuals. It indicates how strong the selective pressure is on the population.
The Breeder's Equation is used to calculate the response to selection (R) and is given by:
R= H² X big S
Where:
H² = Broad sense heritability
Big S = Strength of selection
This equation helps predict how much a trait will change in the next generation due to selection.
Little s: Represents the selection coefficient, which measures the relative fitness of a genotype compared to the most fit genotype.
Big S: Represents the strength of selection, which measures the difference in trait values between the population and the reproducing individuals.
Variation in quantitative traits in natural populations can be measured by comparing the survival of individuals that differ in the trait of interest.
Darwin’s finches on the Galapagos Islands showed that natural selection can influence traits like beak size
Survivorship comparison involves comparing the survival of individuals with different traits (such as size or beak length). This can be done by measuring the size of a trait (denoted as z) before and after selective pressure. The formula for measuring selection is:
i= [(mean after selection)-(mean before selection)]/ square root(variance in that trait)
mean after selection= Za
mean before selection= Zb
the variance in the phenotype/trait= Vp
The measure 𝑖 represents the intensity of selection, indicating the directional pressure on a population. If 𝑖 is positive, it suggests directional selection is acting on the population, favoring certain phenotypes.
The selection gradient measures the relationship between the phenotypic value of a trait and the fitness associated with that phenotype. This relationship is estimated by regression analysis and is represented by the slope (beta):
A steeper slope means a stronger relationship between the trait and fitness.
If beta is positive, larger phenotypic values are selected for (directional selection).
If beta is negative, the phenotype is disfavored, indicating negative selection (selection against the trait).
Peter and Rosemary Grant started a research project on the medium ground finch (Geospiza fortis) in 1973. They tracked reproductive success, measured body size, and created genealogies for the finches. The Grants studied heritable variation for traits like beak length and beak depth.
The heritability (h²) of beak length is 0.65, and the heritability of beak depth is 0.9 in the medium ground finch. Both traits show high heritability, meaning they can be passed down to offspring and are influenced by genetic factors.
Beak size influences the type of seed the finch can easily handle. Larger beaks are suited for cracking hard seeds from caltrop, while smaller beaks are suited for cracking soft seeds from spurge plants.
In 1977, a drought on Daphne Major caused most of the spurge seeds (small, soft seeds) to be unavailable. The average beak depth increased from 9.2 mm before the drought to 9.7 mm afterward, as birds with larger beaks were better able to handle the larger, harder caltrop seeds. This is an example of directional selection.
After the 1977 drought, beak depth in medium ground finches increased, as birds with deeper beaks were more capable of handling the larger, harder seeds that became more abundant during the dry season. Beak depth was found to be heritable in the 1978 offspring.
In 1985, heavy rains caused the spurge plant population to increase. As a result, smaller beaked birds were better suited for eating the small soft seeds from spurge. Beak size in the finch population became more narrow, and having a deep beak was no longer advantageous.
Beak length was developmentally correlated with beak depth. Even though beak length was selected against (birds with shorter beaks were favored), beak length still increased because it is developmentally linked to beak depth, which was positively selected for.
Ecological character displacement occurs when competition for resources favors traits that reduce overlap between species. In the case of the medium ground finch (G. fortis), the large-beaked finch (G. magnirostris) outcompeted them for large seeds. As a result, the mean beak size in G. fortis decreased over time, and smaller beaked individuals were favored. This is an example of ecological character displacement.
The study of Oldfield mice in the Gulf Coast supports the theory of evolution by natural selection. The light and dark mice match the backgrounds they inhabit, which suggests that natural selection through visual predation is shaping the color traits of these mice.
Researchers used plasticine models of light and dark Oldfield mice to test the effect of visual predators. They placed the models on soil backgrounds that either matched or didn’t match the color of the model mice. They found that mismatched models were attacked more frequently by predators than the matching models.
Researchers found that mismatched models of Oldfield mice (those not matching the background color) were attacked much more often by predators. The proportion of attacks on mismatched mice was higher compared to the matching mice, indicating that crypsis (color matching with the environment) offers a survival advantage.
On the Atlantic Coast of Florida and the Gulf Coast, there are populations of light and dark Oldfield mice that match the colors of their respective environments. This indicates directional selection for color matching to avoid predation.
Molecular analysis revealed that the light-colored Oldfield mice from the Gulf Coast and Atlantic Coast are more closely related to the dark-colored mice from the same regions, suggesting two independent origins of the light color morph. This supports the idea of independent evolution of the light color due to local predation pressures and crypsis.
Directional selection is observed in Oldfield mice because mice with colors matching their backgrounds are less likely to be seen and eaten by predators. Over time, mice with advantageous color traits (light or dark, depending on the environment) are selected for, leading to a change in the population's phenotype toward the more advantageous color variant.
Disruptive selection is observed in Gall fly populations because different selective pressures (predators) push the gall size in opposite directions. Large galls are easier for birds to predate, while small galls are more vulnerable to parasitic wasps. As a result, both extremes are favored by selection, leading to two distinct populations.
Birds prefer large galls → easier to access larvae.
Wasps prefer small galls → better for laying eggs.
This results in disruptive selection favoring both extremes.
When both are present:
Stabilizing selection occurs (favors intermediate sizes).
If parasitism is stronger: Directional selection favors smaller galls.
Armor (bony plates/spines) helps protect against large predatory fish.
In freshwater, armor is reduced due to lower predation and nutrient deficiencies.
Lower predation pressure in lakes (fewer large predators).
Nutrient deficiencies in freshwater lakes (low ion concentrations).
The low-eda allele is responsible.
Recessive allele more common in freshwater.
In marine, the allele is rare.
Armor reduction due to:
Lower predation in freshwater.
Nutrient deficiency (low ions) in freshwater lakes.
Results in evolution of armor reduction.
Lactase persistence: Ability to digest milk into adulthood.
Caused by mutations in the LCT gene.
Present in 30% of adults worldwide.
Mutations allow individuals to digest milk sugars (lactose).
Domestication of cattle led to milk consumption.
Milk is nutrient-rich → those who could digest it had a survival advantage.
Lactase persistence provides a nutritional advantage in environments with domesticated animals.
Selective sweep: Strong selection for a beneficial allele (lactase persistence) causes it to become fixed in a population.
Loci near LCT gene are inherited together in those with lactase persistence.
Hitchhiking: The process where nearby alleles increase in frequency due to the selection of a favorable allele (e.g., lactase persistence).
Alleles near the LCT gene are more likely to be inherited together due to their physical proximity on the chromosome.
Molecular fingerprint left by past selective sweeps can be observed.
Independent mutations led to lactase persistence in European and African populations.
Genetic evidence supports independent selective sweeps in each population.
Humans influence selection on many species, both prokaryotes and eukaryotes.
Artificial selection: Humans select for traits in plants and animals that are useful or interesting.
Inbreeding in purebred lines can accidentally select for deleterious recessive alleles.
Cruciferous vegetables (e.g., cabbage, broccoli, cauliflower) are the same species but selected for different traits.
Domesticated crops, like corn (big kernels) and tomatoes (large fruit), have been selected over centuries to be much different from their wild ancestors.
Hunting creates selective pressures by favoring traits that allow animals to survive.
Elephants with tusks are targeted by poaching, so tuskless elephants have higher survival and reproductive success.
This has led to mutations causing tuskless elephants to become more common.
Fishing has caused species like Atlantic cod to undergo directional selection, favoring earlier maturity and smaller body sizes.
Larger fish are often selected against due to fishing practices, leading to declining body size in populations with high fishing pressure.
Advantageous allele starts rare.
A new advantageous allele is unlikely to become fixed unless it increases in frequency when rare.
Example: Darwin's finches and beak size.
Keeps genetic diversity at levels higher than expected by chance.
Occurs due to negative frequency-dependent selection or heterozygote advantage.
Heterozygotes have higher fitness than both homozygotes.
Leads to stable polymorphism where both alleles, A1 and A2, are maintained in the population.
Study of continuous traits and their evolutionary mechanisms.
It measures the difference between individual measurements and the mean.
It assumes population variance, but it's based on sample data since it's not feasible to measure all individuals.
Evolution driven by competition for resources.
Traits shared between species evolve to reduce resource overlap (e.g., beak size in birds).
Camouflage; animals blend with their background to avoid predators.
Sequences derived from a common ancestral gene.
Loci on either side of an allele increase in frequency without selection.
Orthologs: Homologous sequences in different species, separated by speciation.
Paralogs: Homologous sequences within the same species, separated by gene duplication events.
the reason for differentiating is that the two copies can have separate evolutionary histories, many not have the same origin.
Neutral mutations accumulate at a constant rate, correlating to differences between taxa.
Helps in tree reconstruction and calibration of evolutionary timelines.
Caveat: Less accurate for long-diverged taxa.
Molecular data helps trace back traits, even when morphological data is unavailable.
Morphological data is more informative for identifying speciation events.
Coalescence is the merging of homologous gene copies from different individuals to a common ancestor when traced back through generations.
It doesn't imply a single ancestral individual but rather a common ancestor for those alleles.
Coalescence refers to the coming together of homologous sequences tracing back in time.
Convergence is the independent evolution of similar traits or sequences in unrelated lineages.
Homologous alleles are derived from a common ancestral sequence and have descended from the same original genetic source.
Tracing alleles back in time shows that they may not be passed down directly through generations.
Mutations lead to the emergence of new alleles that eventually coalesce into one of many fixed alleles in the population.
Polymorphisms like G → T in BRCA1 can be traced back to see when the mutation occurred.
Sequences with ancestral G and derived T are homologous and coalesce at their common ancestral sequence.
In a population, multiple copies of a sequence can be traced back in time to a common ancestor.
Coalescence occurs when two copies of a sequence eventually trace back to a single common ancestral copy.
Mutations can occur without speciation.
Genes may not change during speciation, leading to different patterns in gene and species trees.
Example: Human genes have different coalescent times; mitochondrial DNA evolves rapidly, while MX1* evolves slowly.
Mitochondrial DNA has a rapid coalescence time (<0.5 million years), reflecting fast evolution.
MX1 gene* has a slow coalescence time (~8 million years), indicating slow evolution.
It occurs when a genetic polymorphism persists through multiple speciation events.
This can cause a different gene tree than the actual species tree, creating confusion in genetic relationships.
For example, humans are more closely related to chimps than gorillas.
Incomplete sorting may cause some parts of the human genome to be more closely related to gorillas than chimps, leading to incorrect conclusions about species relationships.
All regions of the human genome would be more similar to chimps than to gorillas or other species.
No incomplete sorting means genetic differences reflect true species relationships.
Incomplete sorting can cause some genomic regions to appear more closely related to the wrong species, which may produce incorrect phylogenetic trees.
Solution: More data is needed to accurately reconstruct the true evolutionary history.
Single gene analysis can produce conflicting results.
Example: 70% of genetic analyses show humans and chimps are more related, but 30% show humans and gorillas are more related, due to using different genes.
Unambiguous identification of character states (A, T, C, G).
Can use thousands of genetic characters for analysis.
Restricted to extant taxa (living species) or recently living.
DNA rarely survives for over 100k years, making ancient DNA analysis difficult.
Harder to distinguish homoplasy (independent mutations) from homology (shared ancestry).
Morphological data is still needed to support molecular data.
Max Parsimony minimizes the number of evolutionary changes required to explain the data.
Exons are given more weight than introns because they are less likely to undergo homoplasy.
Bootstrapping involves randomly selecting subsets of characters, redoing the tree with those subsets, and then analyzing the consistency of the results.
If the same relationships appear consistently, the tree has high bootstrap support.
Morphological data is used to support molecular data, especially when distinguishing homoplasy from homology.
Helps ensure the reliability of phylogenetic relationships.
Neighbour Joining calculates the distance between taxa based on similarities.
Starts with the most closely related taxa.
Quick but less reliable than other methods.
Can use amino acid sequence differences.
Advantage: Fast, easy for computers to run.
Disadvantage: Less reliable compared to other methods because it doesn't account for ancestral/derived states of characters.
ML uses prior knowledge of how mutations occur in DNA.
Model selection is based on known mutation patterns (e.g., mutations at 3rd position of codon are more likely).
For each tree, ML calculates the probability of observing the data given the tree, and assigns a likelihood value to the tree.
Requires significant computing power.
Similar to Maximum Likelihood, but uses inverse probability.
Starts with a tree shape and calculates the probability, then tweaks it to maximize the probability.
Requires significant computing power and time.
Both require prior knowledge of mutation models.
ML calculates probability of observing the data given the tree.
Bayesian method starts with a tree and tweaks it to maximize the probability of the tree fitting the data.
Both methods are computationally intensive.
ML and Bayesian methods require enormous computational power (days/weeks).
Quick methods like NJ or MP are often run first for initial results.
If multiple methods agree, it gives strong support for a tree.
Polymorphism: A common difference in sequence between related species or individuals of the same species.
Polymorphisms (genetic differences) are often non-selective, meaning they can arise without natural selection acting on them, complicating tree construction.
Pros:
Unambiguous character states (e.g., A, T, C, G)
Can use 100,000s of characters.
Cons:
Restricted to extant taxa (living species)
Need to distinguish between homology and homoplasy (similar traits that are not due to common ancestry).
Fossils suggest tetrapods evolved from lobe-finned fish ancestors.
Molecular phylogeny using 251 genes agrees with morphological phylogeny, showing tetrapods' closest living relatives are coelacanths and lungfish.
The "Out of Africa" hypothesis suggests humans originated in Africa.
Molecular phylogenetic analysis shows the greatest genetic variation is found in native African populations, supporting this hypothesis.
HIV is polyphyletic at two levels:
HIV-1 comes from chimpanzee simian immunodeficiency virus (SIV).
HIV-2 comes from sooty mangabey SIV.
HIV-1 jumped into humans twice, once directly from chimpanzees and once from gorillas.
The virus likely entered humans through bushmeat trade, and subtypes can be traced via molecular phylogeny to specific ape/monkey species.
Moto Kimura.
Much of the variation in genomes is due to genetic drift rather than natural selection.
Mutation accumulation rate is constant and predictable.
Mutations accumulate faster in regions of the genome that don’t affect phenotype.
Synonymous mutations accumulate faster than non-synonymous mutations.
It serves as the null hypothesis for natural selection, proposing that many genetic changes are due to drift.
neutral mutations should accumulate at a roughly constant rate over time.
A 1970 study comparing cytochrome c gene differences between pairs of mammalian species
A positive linear relationship was observed between the number of mutations and time since species divergence.
Calibration (e.g., via fossils or geographic events) provides a time start point for measuring evolutionary time.
Without it, you can measure time passed, but not when an event started
Reversals or mutations that revert back to the original base pair can occur.
For closely related species, the slope of differentiation is high, but as divergence time increases, the slope plateaus due to saturation of mutations.
Saturation occurs when new mutations overwrite older ones, causing a plateau in sequence differences after ~10-20 million years.
No, rates of sequence evolution are not constant across lineages, as factors like generation time and metabolic rate influence divergence rates.
Sequence divergence rates are negatively correlated with body size (and generation time); endotherms evolve faster than ectotherms.
When generation times are fast, such as with viruses or bacteria, allowing for recent event dating (e.g., HIV evolution).
By using preserved blood samples and molecular clocks, researchers estimated HIV’s jump to humans occurred between 1883-1925.
Increased bushmeat trade during colonization likely facilitated the jump of HIV from chimps to humans.
Neutral theory suggests that most genetic variation in a population is due to genetic drift, not natural selection. It acts as the null hypothesis for detecting natural selection.
Selective sweeps leave molecular evidence of past natural selection through hitchhiking sequences. This evidence typically persists for around 10,000 years
Researchers compare synonymous mutations (neutral) and non-synonymous mutations (adaptive). If non-synonymous mutations accumulate faster than synonymous ones, it suggests directional selection.
In stabilizing selection, there are more synonymous mutations than non-synonymous mutations, as stabilizing selection conserves the gene's function.
Weak selection pressures may not leave a noticeable trace, so strong selection is necessary to detect footprints of selection in genetic data.
Prokaryotes (bacteria) have streamlined genomes with most of the DNA coding for proteins, while eukaryotes often have larger, more complex genomes, including non-coding regions like pseudogenes.
Some prokaryotes, especially intracellular endosymbionts, have small genomes because they rely on their host for many functions and have abandoned many genes.
Eukaryotes often have pseudogenes, parasitic DNA (e.g., viral remnants), and other non-coding regions, which contribute to the larger, more complex genome
The time it takes for a cell to divide is positively related to genome size, as more DNA requires more time to replicate during cell division.
Organisms with smaller genomes may have reduced DNA because they rely on a host organism for some biological functions, often seen in endosymbionts.